Journal of the Lepidopterists Society 53(2),1999, 65-71 PRELIMINARY ESTIMATES OF LEPIDOPTERA DIVERSITY FROM SPECIFIC SITES IN THE NEOTROPICS USING COMPLEMENTARITY AND SPECIES RICHNESS ESTIMATORS MICHAE L G. POGUE Systematic Entomology Laboratory, P. S. I. , Agricultural Research Service, U.S. D epartment of Agriculture, % D ep artment of Entomology, National Museum of Natura! HistOIY, Smithsonian Institution, Washington, D.C. 20560-0168, USA ABSTRACT. Lepidoptera were collected and species richness and compl ementarity or uniqueness were compared between two rainfor­ est sites: Pakitza, Peru and Beni , Bolivia. The total number of species collected from hoth sites was 1,879 of which 60 were shared resulting in a complementarity of96.8%. Non-parametric equations and species accumulation curves of Hemiceras Guenee (Lepidoptera: Notodontidae) were used to compare species richness between three rainforest sites, Pakitza and Tambopata, Peru and Reserva Ethnica Waorani, Onkone Gare, Ecuador. Cluster analysis, using complementarity values for selected sites was used to determine altitudinal relationships between sites in Costa Rica; relationships between forest types in Brazil; and faunal differences among sites in western Amazonia using Hemiceras. Additional key words: Biodiversity, Notodontidae, Hemiceras, Chao, jackk'l1ife . Biodiversity as defined by E. O. Wilson (Reaka­ taxa around the globe. After a site has been sampled, Kudla et al. 1997) is "everything". Biodiversity encom­ species richness estimates are used to predict how passes the genes within a Single local population or many species were missed during the sampling pro­ species, the species within a local community, and cess, thus arriving at an estimated number of species communities comprising the diverse ecosytems of the based on the actual number observed plus the number world. Life on earth is supported by the interactions missed. By using species lists, either generated by and products produced by all other life on earth. With­ sampling at a site or from museum collections, species out biodiversity there would be no life on earth as we composition among sites can be compared. These know it. Therefore, it is essential that biologists begin comparisons can then be used in setting policy and to document and record biodiversity, whether it be making informed conservation and management deci­ how many species of insects are in your backyard to sions. how many species of trees in a forest to how many for­ The goals of this paper are 1) to emphaSize the im­ est types in the world. portance of adequate sampling, 2) to assess whether Anyone with an interest in natural history can begin inadequate sampling can still be useful in predicting to study their local biodiversity and to document it. species richness and complementarity between study One basic element of biodiversity is to know how sites, and 3) to prOvide a method of using complemen­ many species are present at a particular site. This pa­ tarity to compare faunal relationships between sites. per will outline how anyone can begin to document lo­ To accomplish these goals, I compared overall species cal biodiversity by gathering data on species richness richness and complementarity of Lepidoptera from or the number of species present at a site at a particu­ two rainforest sites, one in Peru and the other in Bo­ lar point in time. KnOwing what species are present at livia. In addition, I estimated species richness from a site is essential because it is the first step in under­ species accumulation curves and non-parametric esti­ standing the interactions between the species docu­ mators to compare the Hemiceras Guenee (Notodon­ mented and their interactions within the local commu­ tidae) fauna between 2 sites in Peru and 1 in Ecuador. nity and ecosystem. Finally, museum specimens of Hemiceras were used Presently, no one has much of an idea exactly how for faunal comparisons among sites in Costa Rica, many species are present today on earth. Estimates of Brazil, and western Amazonia, using complementarity the number of species worldwide range from 3 to 100 and cluster analysis. million (Erwin 1982, 1983, Stork 1988, Hodkinson & Casson 1991, May 1992, Raven and Wilson 1992). The MATERIALS AND M E THODS study of species richness and complementarity, or how Lepidoptera complementarity in SW Amazo­ different species composition is between sites, is es­ nia. Lepidoptera were collected at two rainforest sites: sential to assessing global biodiversity patterns. Beni, Bolivia and Pakitza, Peru. The Beni study site at To address the question of world insect diversity 14°49' S, 66°28'W is 40 km E of San Borja, at 250 me­ one must get accurate estimates of Site-specific species ters elevation. Pakitza is located on the Rio Manu at richness for a variety of taxa (Colwell & Coddington 11 °56'4TS, 71°1 TOO'W within the large drainage 1994), and then to compare these species lists to mea­ basin of the Rio Madre de Dios in southeastern Peru, sure relative levels of overlap and richness of these at 356 m elevation, approximately 550 km NW of Beni. 66 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY 40 35 30 25 <f) Q) 'u Q) j5- 20 0 z0 15 10 -+- Pakitza ;J ____ Tambopata ~ Onkone Gare 5 Trap Nights FIG. 1. Species accumulation curves of Hemiceras at Pakitza and Tambopata, Peru, and Onkone Care, Ecuador. Samples were collected between August 26-Septem­ The complementarity between the two sites is the ber 15, 1987 (Beni) and from September 27-0ctober proportion of the unique species to the pooled rich­ 5, 1987 (Pakitza). Adult moths were collected by UV ness, or light traps, spread, identified to family, sorted to mor­ pho-species, and counted. Voucher specimens have (3) been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D .C. Collecting effort was defined as the number of t~\lP ' Hemiceras species richness in western Amazo­ nights: 12 and 8 trap nights in Pakitza and Beni, re­ nia. The genus Hemiceras (Lepidoptera: Notodonti­ spectively. Other studies have included the number of dae), representing 245 species, was used as an indica­ person-hours spent collecting (Coddington et al. 1991, tor group for estimating species richness at three Robbins et al. 1996), number of collecting days (Lou­ rainforest sites: Pakitza and Rio Tambopata Research ton et al. 1996), or trap nights (approximately 12 hours Station in southeastern Peru, and Onkone Care, in in length). Ecuador. The Lepidoptera faunas of Pakitza and Beni were Rio Tambopata Research Station, at 14°14'S, compared using complementarity (Colwell and Cod­ 69°11'W, is located on the RIO Tambopata, 30 air km dington 1994). In comparing two sites, j and k, the SE of Puerto Maldonado, Madre de Dios, Peru, at first site has a species richness of Sj and the second 290 m elevation. Onkone Care, at 00038'S, 76°36W, is site has Sk If the number of species in common be­ a research station within the Reserva Ethnica Waorani, tween both sites is Vjk, then the total species richness Ecuador, at 220 m elevation. Tambopata had a total of for both sites is 29 trap nights from November 2-25, 1979 and Sep­ tember 16-21, 1990. The 11 trap nights at Onkone (1) Care were January 10, 12,13-18,25; June 20; and July 16,1994. and the number of species unique to both sites (Uik ) is Species accumulation curves (Fig. 1) were used to plot the cumulative number of new species collected (2) over unit effort (number of trap nights) at each of the VOLUME 53, N UMBER 2 67 FTC. 2. Localiti es of Costa Rican sites used in cluster analysis of dissimilarity matrix in Table 3. FIG. 3. Localities of South American sites used in cluster analyses of dissimilarity matrices in Tables 4 and 5. three sites. To extrapolate total species richness from jackknife estimator of species richness is based on the the species accumulation curves at each site, four dif­ number of species that occur in only one sample L, ferent nonparametric equations were compared: 1) Chao 1, 2) Chao 2, 3) first-order jackknife, and 4) sec­ 1 jackknife = Sobs + L(n - lin), (6) ond-order jackknife. Chao (1984) developed a simple estimator of the where n is the number of samples. true number of species at a given site based on the The second-order jackknife (Burnham & Overton number of rare species in the pooled sample j. This is 1978,1979) is like the Chao 2 estimator where L is the considered an abundance-based estimator because it is number of species that occur in only one sample and based on the number of species that are only repre­ M is the number of species that occur in exactly two sented by only 1 or 2 individuals to estimate overall samples, species richness. Colwell and Coddington (1994) called this Chao 1, 2 jackknife = Sobs + 1/2 [L(2n - 3)/n - M(n - 2)2/n(n - 1)], Chao 1 = Sobs + a2/2b, (4) (7) where Sobs is the observed number of species in a sam­ where n is the number of samples. ple, a is the number of species that are only repre­ Hemiceras faunal comparisons between three sented by one specimen in the pooled sample (Single­ tropical regions. SpeCies lists of Hemiceras were com­ tons), and b is the number of species represented by piled from specimens in the National Museum of Natu­ two specimens in the pooled sample (doubletons). ral History, Smithsonian Institution, Washington, D.C., This estimator works well when the samples contain a and were used to examine faunal relationships among large number of rare species (Chao 1984), which fre­ sites in Costa Rica, Brazil, and western Amazonia. In quently occurs when sampling diverse groups such as Costa Rica, six sites were chosen to see how altitude af­ insects.